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Hormone-Induced Spawning of Cultured Tropical Finfishes

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Hormone-Induced Spawning of Cultured Tropical Finfishes ADVANCES IN TROPICAL AQUACULTURE. Tahiti, Feb. 20 - March 4 1989 AQUACOP 1FREMER Acres de Colloque 9 pp. 519 F39 49 Hormone-induced spawning of cultured tropical finfishes C.L. MARTE Southeast Asian Fisheries Development Genie. Aquaculture Department, Tig- bauan, ILOILO, Philippines Abstract — Commercially important tropical freshwater and marine finfishes are commonly spawned with pituitary homogenate, human chorionic gonadotropin (HCG) and semi-purified fish gonadotropins. These preparations are often adminis- tered in two doses, a lower priming dose followed a few hours later by a higher resolving dose. Interval between the first and second injections may vary from 3 - 24 hours depending on the species. Variable doses are used even for the same species and may be due to variable potencies of the gonadotropin preparations. Synthetic analogues of luteinizing hormone-releasing hormone (LHRHa) are becoming widely used for inducing ovulation and spawning in a variety of teleosts. For marine species such as milkfish, mullet, sea bass, and rabbitfish, a single LHRHa injection or pellet implant appears to be effective. Multiple spawnings of sea bass have also been obtained following a single injection or pellet implant of a high dose of LHRHa. In a number of freshwater fishes such as the cyprinids, LHRHa alone however has limited efficacy. Standardized methods using LHRHa together with the dopamine antagonists pimozide, domperidone and reserpine have been developed for various species of carps. The technique may also be applicable for spawning marine teleosts that may not respond to LHRHa alone or where a high dose of the peptide is required. Although natural spawning is the preferred method for breeding cultivated fish, induced spawning may be necessary to control timing and synchrony of egg production for practical reasons. INTRODUCTION There are more than 80 species of freshwater and marine fish cultured in Asia. Of these, about 50 species are cultured in tropical South and Southeast Asia (Rabanal, 1988). For most of the cultured species, wild fry from natural sources is insufficient to supply the requirements for culture. Moreover, the fry supply is dependent on the season, and fluctuates with environmental and climatic conditions. Overfishing, pollution and various human activities have caused the destruction of natural spawning and fry grounds contributing largely to 520 C.L. Marte the reduction in fry catch. Of the numerous species under cultivation only a few are bred in captivity. For those species that spontaneously breed under captive condition, the time of spawning is often not predictable. Problems such as viability of naturally spawned eggs and technical difficulties in egg and larvae collection are constraints to mass-scale fry production. Most of the information on the physiological processes involved in the hormonal control of fish reproduction are derived from studies on a few species notably the salmonids, goldfish, and common carp. Although the basic biological principles arising from studies on these fish apply to tropical species, the diversity of fish cultured in the tropics present numerous problems. This paper summarizes the recent developments in induced breeding of impo rtant species cultured in Southeast Asia. PRACTISES IN INDUCED BREEDING Induced breeding of captive fish may be approached in two ways, hormonal and environmental. For most of the tropical cultured fish, the specific environmental cues that trigger ovulation and spawning have not been identified. Asian fish breeders however have succesfully developed methods that stimulate spawning conditions in a few freshwater species. Environmental manipulation to induce ovulation and spawning in fish has been reviewed by Lam (1983) and Lam and Munro (1987). Specific examples of tropical fish where environmental cues are known and used to stimulate breeding activities are cited in Lam (1985). Induction of spawning using hormones provides a direct control over the final stages of the reproductive cycle in teleosts. Hormonal induction of spawning has been the subject of many recent reviews (Harvey and Hoar, 1979; Lam, 1982, 1985; Donaldson and Hunter, 1983; Crim et al., 1987; Abraham, 1988). The physiological mechanisms involved in the final stages of oocyte maturation, ovulation and egg release have been tho- roughly reviewed (Fostier and Jalabert, 1982; Goetz, 1983). Harvey and Hoar (1979) and Davy and Chouinard (1980) discuss traditional practises followed in induced spawning of tropical fish. Ovulation and spawning in teleosts as in other vertebrates are controlled by several interacting factors. Environmental stimuli are transla- ted by the brain into neural signals which result in the release of gonadotropin releasing hormone (GnRH) and/or inhibition of the release of gonadotropin release inhibiting factor (GnRIF) causing the pituitary to secrete gonadotropins (GtH) (Peter, 1982; 1983a; Peter et al., 1986; Lin and Peter, 1986). When a ce rtain GtH level is reached, vitellogenic oocytes undergo the process of final oocyte maturation : the germinal vesicle migrates to the periphery; theca and granulosa cells of the follicle are stimulated to secrete a maturation-inducing steroid (MIS); and the MIS induces germinal vesicle breakdown (GVBD) (Nagahama, 1983; Fostier and Jalabert, 1983; Goetz, 1983). Evidence from studies on goldfish, salmonids and other species point to 17-alpha, 20-beta progesterone (17a, 2013P) as the MIS (Nagahama,1983; Scott and Canario, 1987) although other related progestogens have been 49 - Hormone-induced spawning of cultured tropical finfishes 521 identified (Scott and Canario, 1987). Corticosteroids have also been implicated (Goetz, 1983; Fostier and Jalabert, 1983) but may play only a supportive role (Jalabert, 1976). An additional role for 17a, 20(3P and other progestogens as reproductive pheromones particularly in those species where very high levels are found, has been suggested (Scott and Canario, 1987). Hypophysation and Human chorionic gonadotropin (HCG) administration. Traditional methods of induced breeding involve the injection either intramuscularly or intraperitoneally of crude pituitary extracts. The pituitary extract is usually administered to the female in two doses, a stimulating dose followed after a variable time interval by a second resolving dose. Fresh or preserved pituitary glands from mature fish of the same species (homoplastic) or from other, usually related species (hete- roplastic) are used. In some cases, glands from immature fish have been used but higher doses are required (Harvey and Hoar, 1979). Females that do not respond after the second injection are injected a third or even more doses. Injecting multiple doses however has seldom been successful and females regress probably as a result of stress from excessive handling. Males are injected the same or half the dose given to the female usually at the time the second or resolving dose is administered. Doses are given as fresh or dry weights of pituitary gland per unit body weight of the broodfish or in dose units, defined as the ratio of the body weight of the donor and the body weight of the recipient. Standardization of hypophysa- tion is difficult since the potency of the pituitary extract depends on the age, sex and state of maturity of the donor. Method of collection and the technique used to preserve the pituitary also vary. Species specificity of gonadotropins has been demonstrated (Fontaine et al, 1972; Varikul and Sritongsook, 1981) and is an impo rtant factor to consider. The supply of pituitary glands is a problem and although crude or partially purified pituitary extracts with assayed gonadotropin potency is commercially available, the cost for the-Asian fish breeder is prohibitive. The problems of standardization and cost of hormone preparations are partly solved with the use of mammalian gonadotropin preparations. Two are available in purified form, human chorionic gonadotropin (HCG) and pregnant male serum (PMS). The dosage used varies widely between species and may be related to how closely HCG and PMS resemble the endogenous gonadotropin in each species (Lam, 1982). HCG has been successfully used in most species bred in Southeast Asia (examples are given in the succeeding sections). HCG is available and convenient to use although still expensive. The possibility that injecting HCG and pituitary extracts for several consecutive years to the same broodstock may result in the development of an immune response has been pointed out (Lam, 1982; Billard et al 1987). Luteinizing hormone-releasing hormone (LHP/-l} LHRH, a hypothalamic decapeptide and its synthetic analogues have been shown to stimulate gonadotropin secretion in teleosts (Crim et al., 1987; Peter 1983a and b; Lin and Peter, 1986). The effectiveness of LHRH analogues in inducing ovulation and spawning of cultured fish was first 522 C.L. Marte demonstrated in various species of carps by Chinese researchers (Anon 1977). The practise however was not widely adopted because consistent results were not obtained. Recent studies have demonstrated the presence of a gonadotropin- release inhibiting factor in goldfish. Further evidences identify GRIF to be the catecholamine dopamine (for review, see Peter et aL, 1986). GRIF's inhibitory effect on GtH release is blocked by administration of dopamine receptor antagonists such as pimozide or metoclopramide (Chang et al., 1984; Sokolowska et aL,1984, 1985; Peter et al., 1985). Administration of dopamine antagonists potentiates LHRHa
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